Peculiarities of Activation of the Upper Limb Muscles in Realization of Two-Joint Movements in Humans

In tests on humans, we recorded EMG activity from the muscles flexing and extending the forearm and shoulder in the course of realization of sequential single-joint and simultaneous two-joint movements of the upper limb. As was shown, the shoulder muscles m. biceps brachii and m. triceps brachii are involved in flexion/extension of both elbow and shoulder joints. Central commands sent to the above muscles in the course of a two-joint movement could be considered a superposition of the central commands coming to the same muscles in realization of the corresponding sequential single-joint movements with the same changes in the angles of the elbow and shoulder joints. External loadings applied in the direction of extension of the elbow and shoulder joints induced, in general, similar changes in coordination of the activity of muscles moving the forearm and shoulder under conditions of both single-joint and two-joint movements. These facts allow us to suppose that coordination of the muscle activity in two-joint movements depends to a greater extent on the forces influencing limb links than on the mode of realization of the movements (two sequential single-joint movements vs a two-joint movement corresponding to the above motor events).     

Activation of the Shoulder-Belt and Shoulder Muscles in Two-Joint Arm Movements Performed in Humans with the Action of Opposite Loadings

In tests on four volunteers, we examined coordination of central motor commands (CMCs) controlling slow two-joint movements of the arm within the horizontal plane. Current amplitudes of EMGs recorded from six muscles of the shoulder belt and shoulder and subjected to full-wave rectifying and low-frequency filtration were considered correlates of these commands. In particular, we studied the dependence of coordination of CMCs on the direction of an external force applied to the distal forearm part. As was found, coordination of CMCs significantly depends on the direction of the force flexing the elbow joint. According to our observations, EMGs of definite muscles in the case of performance of a two-joint movement can, in a first approximation, be presented as linear combinations of the EMGs recorded in the course of separate sequential single-joint movements under conditions of shifting the reference point of the hand toward the same point of the operational space as that in the two-joint movement. These data can be interpreted as confirmation of the principle of superposition of elementary CMCs in the performance of complex movements of the extremity.     

Activation of the Shoulder Belt and Shoulder Muscles of Humans Related to Different Rates of Generation of Two-Joint Efforts by the Forearm

We studied coordination of central motor commands (CMCs) coming to the muscles that flex and extend the shoulder and elbow joints in the course of generation of voluntary isometric efforts of different directions by the forearm. Dependences of the characteristics of these commands on the direction of the effort and rate of its generation were analyzed. Amplitudes of rectified and averaged EMGs recorded from a number of shoulder belt and shoulder muscles were considered correlates of the CMC intensity. The development of the effort of a given direction and rate of rise was realized in the horizontal-plane operational space; the arm position corresponded to the 30 deg angle in the shoulder joint (external angle with respect to the frontal plane) and 90 deg angle in the elbow joint. We plotted sector diagrams of the relative changes in the level of dynamic and stationary phases of EMG activity of the studied muscles for the entire set of directions of the efforts generated with different rates of rise. In the course of formation of rapid two-joint isometric efforts, realization of nonsynergic motor tasks (extension of one joint and flexion of another one, and vice versa) required significant activation of muscles of different functional directions for both joints. Time organization of EMG activity of extensors and flexors of the shoulder and elbow joints related to the maximum and relatively rapid generation of the effort (rise time 0.12 to 0.13 and 0.25 sec, respectively) was rather complex and included dynamic and stationary phases. With these time parameters of generation of the efforts (both flexion and extension), the appearance at the stationary effort of 40 N was controlled based on coordinated interaction of dynamic phases of the activation of agonistic and antagonistic muscles. It is concluded that CMCs coming to extensors and flexors of both joints upon generation of rapid isometric efforts are rather similar in their parameters to those under conditions of realization of the forearm movements in the space in an isotonic mode.     

Peculiarities of Activation of the Shoulder Belt and Shoulder Muscles in Generation of Different-Direction Isometric Efforts by the Forearm

We studied central motor commands, CMCs, coming to the muscles that flex and extend the shoulder and elbow joints in the course of generation of voluntary isometric efforts of different directions by the forearm; the efforts were initiated according to a visual signal. Amplitudes of EMGs recorded from the muscles of the shoulder belt and shoulder and subjected to full-wave rectification and low-frequency filtration were considered correlates of the CMC intensity. An effort of the preset direction was developed within the operational space of the horizontal plane with angles 30 deg in the shoulder joint (external angle with respect to the frontal plane) and 90 deg in the elbow joint. We plotted sector diagrams of the logarithmic coefficient of the intensity increment of EMGs of the above muscles for the entire set of directions of generated efforts with a 15- or 20-deg step. Orientations of the maxima of EMG activity of the given muscles were rather close to the directions of the maxima of the force moments generated by these muscles. In most cases, a shift of the direction by one gradation with respect to the EMG maximum in the respective muscle resulted in a significant decrease in the level of EMG activity. It is shown that preferential activation of the muscles agonistic with respect to the examined direction of the generated effort was, as a rule, accompanied by coactivation of the antagonist muscles. When “two-joint” isometric efforts are formed, realization of the socalled synergic muscle tasks (where prevailing contractions of the muscles of the same functional direction for both joints coincide, i.e., flexion-flexion or extension-extension) is organized in a simpler manner. The programs of “nonsynergic” contractions (flexion of one joint and extension of another one, or vice versa) are more complex. In different subjects, considerably dissimilar patterns of EMG activity in muscles influencing these joints could be observed.     

Coupling between muscle activities and muscle torques during horizontal-planar arm movements with direction reversal.

In this study we investigated the hypothesis that the simple set of rules used to explain the modulation of muscle activities during single-joint movements could also be applied for reversal movements of the shoulder and elbow joints. The muscle torques of both joints were characterized by a triphasic impulse. The first impulse of each joint accelerated the limb to the target and was generated by an initial burst of the muscles activated first (primary mover). The second impulse decelerated the limb to the target, reversed movement direction and accelerated the limb back to the initial position, and was generated by an initial burst of the muscles activated second (secondary movers). A third impulse, in each joint, decelerated the limb to the initial position due to the generation of a second burst of the primary movers. The first burst of the primary mover decreased abruptly, and the latency between the activation of the primary and secondary movers varied in proportion with target distances for the elbow, but not for the shoulder muscles. All impulses and bursts increased with target distances and were well coupled. Therefore, as predicted, the bursts of muscle activities were modulated to generate the appropriate level of muscle torque.     

Velocity-Based Planning of Rapid Elbow Movements Expands the Control Scheme of the Equilibrium Point Hypothesis

According to the equilibrium point hypothesis of voluntary motor control, control action of muscles is not explicitly computed, but rather arises as a consequence of interaction between moving equilibrium position, current kinematics and stiffness of the joint. This approach is attractive as it obviates the need to explicitly specify the forces controlling limb movements. However, many debatable aspects of this hypothesis remain in the manner of specification of the equilibrium point trajectory and muscle activation (or its stiffness), which elicits a restoring force toward the planned equilibrium trajectory. In this study, we expanded the framework of this hypothesis by assuming that the control system uses the velocity measure as the origin of subordinate variables scaling descending commands. The velocity command is translated into muscle control inputs by second order pattern generators, which yield reciprocal command and coactivation commands, and create alternating activation of the antagonistic muscles during movement and coactivation in the post-movement phase, respectively. The velocity command is also integrated to give a position command specifying a moving equilibrium point. This model is purely kinematics-dependent, since the descending commands needed to modulate the visco-elasticity of muscles are implicitly given by simple parametric specifications of the velocity command alone. The simulated movements of fast elbow single-joint movements corresponded well with measured data performed over a wide range of movement distances, in terms of both muscle excitations and kinematics. Our proposal on a synthesis for the equilibrium point approach and velocity command, may offer some insights into the control scheme of the single-joint arm movements.     

A method for estimating three-dimensional human arm movement with two electromagnetic sensors

Measurement of upper-limb movements is important in various domains. In this article, an upper-limb three-dimensional movement recording technique is proposed based on only two electromagnetic sensors. Two joints are considered with a total of seven degrees of freedom (DoF; three translations and four rotations). The chosen sequence of joint rotations is compliant with ISB recommendations: the shoulder is modelled with a ball and socket joint with three DoF and the elbow with a one DoF revolute joint. This article is focused on the procedure used to calibrate and sense the upper-limb movements from the raw data coming from the flock of birds sensors. The principle of the method is to define the centre of the wrist, elbow and shoulder joints in the frame of the adequate sensor. This operation is done by performing calibration gestures. Results are proposed and commented.     

Effects of postural muscle fatigue on the relation between segmental posture and movement.

The purpose of this study was to examine whether fatigue of postural muscles might influence the coordination between segmental posture and movement. Seven healthy adults performed series of fifteen fast wrist flexions and extensions while being instructed to keep a dominant upper limb posture as constant as possible. These series of voluntary movements were performed before and after a fatiguing submaximal isometric elbow flexion, and also with or without the help of an elbow support. Surface EMG from muscles Delto?deus anterior, Biceps brachii, Triceps brachii, Flexor carpi ulnaris, Extensor carpi radialis were recorded simultaneously with wrist, elbow and shoulder accelerations and wrist and elbow displacements. Fatigue was evidenced by a shift of the elbow and shoulder muscles EMG spectra towards low frequencies. Kinematics of wrist movements and corresponding activations of wrist prime-movers, as well as the background of postural muscle activation before wrist movement were not modified. There were only slight changes in timing of postural muscle activations. These data indicate that postural fatigue induced by a low-level isometric contraction has no effect on voluntary movement and requires no dramatic adaptation in postural control.     

A method for estimating three-dimensional human arm movement with two electromagnetic sensors

Measurement of upper-limb movements is important in various domains. In this article, an upper-limb three-dimensional movement recording technique is proposed based on only two electromagnetic sensors. Two joints are considered with a total of seven degrees of freedom (DoF; three translations and four rotations). The chosen sequence of joint rotations is compliant with ISB recommendations: the shoulder is modelled with a ball and socket joint with three DoF and the elbow with a one DoF revolute joint. This article is focused on the procedure used to calibrate and sense the upper-limb movements from the raw data coming from the flock of birds sensors. The principle of the method is to define the centre of the wrist, elbow and shoulder joints in the frame of the adequate sensor. This operation is done by performing calibration gestures. Results are proposed and commented.     

Control of the Elbow Extensor Muscles in Slow Targeted Extensions of the Arm in Humans

Relations between the kinematic parameters of slow (non-ballistic) targeted extension movements in the elbow joint of humans and characteristics of the movement-related EMG activity in the two heads of the m. triceps brachii were analyzed. Test movements were performed under conditions of application of non-inertional external loadings directed toward flexion. It was shown that the movement-related EMG activity of the elbow extensors, similarly to what was observed in the flexors at flexion movements with the same parameters, demonstrates a complex structure and includes dynamic and stationary phases. In the former phase, in turn, initial and main components can be differentiated. The rising edge and decay of the main component of the dynamic extensor EMG phase could be approximated by exponential functions; this component was never split into a few subcomponents. Dependences between the amplitudes of m. triceps brachii EMG phases and the amplitude of the movement (or external loading) were, as a rule, nonlinear but monotonic. An increase in the test movement velocity led to an increase in the rate of rise of the rising edge of the dynamic EMG phase, while an increment in the amplitude was less significant. Under the used test conditions, the activity of the elbow extensors was usually accompanied by some coactivation of the antagonists (m. biceps brachii). It is concluded that motor commands coming to the elbow extensors at performance of the extension test movements differ from motor commands to the flexors at analogous flexion test movements by a simpler structure and more tonic pattern. Biomechanical specificities of fixation of the mentioned muscle groups to the arm bones (stability of the moment for application of the extensor force under conditions of changing the joint angle vs variable moment of the flexor force) are considered one of the main reasons for such specificity of the patterns of the extensor and flexor motor commands.     

Morphology and function of the shoulder joint of bats (Mammalia: Chiroptera)1

The shoulder joint of the Microchiroptera shows a remarkable morphological variation that has been studied in 20 individual bats from 15 species and 11 families. The basic morphology of the shoulder joint, with a globular humeral head and a corresponding glenoid cavity, is found in the Megachiroptera and, within the Microchiroptera, in the Rhinopomatidae. Besides this basic shoulder joint, there are two derived joint types: the derived and specialized shoulder joint with a single articular surface on the scapula and a more-or-less oblong humeral head, and the derived and specialized shoulder joint with secondary articular surfaces on the trochiter and on the dorsal aspect of the scapula. The first type of derived joint is most strikingly developed in the Mormoopidae and the Noctilionidae, the second one in the Vespertilionidae and the Molossidae. It is suggested that both types of derived shoulder joints have the functional significance of reducing the pronatory movements of the abducted forearm during the downstroke of the wing-beat cycle. This suggested function of the secondary shoulder joint is a new approach to understanding this very peculiar structure. In species with these specialized shoulder joints, the downstroke musculature is comparatively better developed and the M. serratus ant. post. div. comparatively less well developed. A hypothesis is offered to explain and combine the osteological and myological findings. Each of the derived types of shoulder joints has developed independently more than once through parallel evolution.     

Mechanical energy costs of human movement: an approach to evaluating the transfer possibilities of two-joint muscles

Different methods of calculating the mechanical energy cost of a movement presented in the literature can give results differing by an order of magnitude. The assumptions made concerning the transfer of energy between different parts of the body are part of the problem. This investigation assesses the role of transfer in energy saving and specifically, the possibility of two-joint muscles reducing the mechanical energy cost of a movement compared to a system having one-joint muscles only. An algorithm was developed which recruited one-joint or both one- and two-joint muscles to supply the net joint moments. The work performed under these two conditions was then compared. It was found that activation of both one- and two-joint musculature reduced the mechanical work cost during walking by between 7 and 29% over that required by single-joint musculature alone. This investigation supports suggestions in the literature that one of the functions of two-joint musculature is to reduce the mechanical energy cost and probably the metabolic cost of movement.     

Mechanical energy expenditure on human movement and the anthropomorphic mechanism]

Mechanical energy expenditures of the man and anthropomorphic locomotion machine during movement are compared theoretically. Sources of the mechanical energy affecting movement of human's lower extremity are modelled by 8 muscles, 3 of which are the two-joint muscles. The model of the lower extremity of anthropomorphic locomotion machine is moved by joint moments. It was shown that in the same movement the model of the human lower extremity can spend less mechanical energy than that of the model of the anthropomorphic locomotion machine. It is caused by the presence of two-joint muscles in the first model. Such an economy of mechanical energy expenditures realized by the two-joint muscle is possible at simultaneous execution of three conditions: 1) signs of the muscle powers, which are produced by that muscle at both joints, are opposite; 2) moments produced by that muscle at each of both joints have the same direction with the joint moments at these joints; 3) one-joint antagonistic muscles are not active. An expression which makes it possible to estimate the mechanical energy savings by the two-joint muscles during humans' movement was developed.     

Interaction torque contributes to planar reaching at slow speed

Background

How the central nervous system (CNS) organizes the joint dynamics for multi-joint movement is a complex problem, because of the passive interaction among segmental movements. Previous studies have demonstrated that the CNS predictively compensates for interaction torque (INT) which is arising from the movement of the adjacent joints. However, most of these studies have mainly examined quick movements, presumably because the current belief is that the effects of INT are not significant at slow speeds. The functional contribution of INT for multijoint movements performed in various speeds is still unclear. The purpose of this study was to examine the contribution of INT to a planer reaching in a wide range of motion speeds for healthy subjects.

Methods

Subjects performed reaching movements toward five targets under three different speed conditions. Joint position data were recorded using a 3-D motion analysis device (50 Hz). Torque components, muscle torque (MUS), interaction torque (INT), gravity torque (G), and net torque (NET) were calculated by solving the dynamic equations for the shoulder and elbow. NET at a joint which produces the joint kinematics will be an algebraic sum of torque components; NET = MUS - G - INT. Dynamic muscle torque (DMUS = MUS-G) was also calculated. Contributions of INT impulse and DMUS impulse to NET impulse were examined.

Results

The relative contribution of INT to NET was not dependent on speed for both joints at every target. INT was additive (same direction) to DMUS at the shoulder joint, while in the elbow DMUS counteracted (opposed to) INT. The trajectory of reach was linear and two-joint movements were coordinated with a specific combination at each target, regardless of motion speed. However, DMUS at the elbow was opposed to the direction of elbow movement, and its magnitude varied from trial to trial in order to compensate for the variability of INT.

Conclusion

Interaction torque was important at slow speeds. Muscle torques at the two joints were not directly related to each other to produce coordinated joint movement during a reach. These results support Bernstein's idea that coordinated movement is not completely determined by motor command in multi-joint motion. Based on the data presented in this study and the work of others, a model for the connection between joint torques (muscle and passive torques including interaction torque) and joint coordination is proposed.     

Analysis of an optimal control model of multi-joint arm movements

 In this paper, we propose a model of biological motor control for generation of goal-directed multi-joint arm movements, and study the formation of muscle control inputs and invariant kinematic features of movements. The model has a hierarchical structure that can determine the control inputs for a set of redundant muscles without any inverse computation. Calculation of motor commands is divided into two stages, each of which performs a transformation of motor commands from one coordinate system to another. At the first level, a central controller in the brain accepts instructions from higher centers, which represent the motor goal in the Cartesian space. The controller computes joint equilibrium trajectories and excitation signals according to a minimum effort criterion. At the second level, a neural network in the spinal cord translates the excitation signals and equilibrium trajectories into control commands to three pairs of antagonist muscles which are redundant for a two-joint arm. No inverse computation is required in the determination of individual muscle commands. The minimum effort controller can produce arm movements whose dynamic and kinematic features are similar to those of voluntary arm movements. For fast movements, the hand approaches a target position along a near-straight path with a smooth bell-shaped velocity. The equilibrium trajectories in X and Y show an ‘N’ shape, but the end-point equilibrium path zigzags around the hand path. Joint movements are not always smooth. Joint reversal is found in movements in some directions. The excitation signals have a triphasic (or biphasic) pulse pattern, which leads to stereotyped triphasic (or biphasic) bursts in muscle control inputs, and a dynamically modulated joint stiffness. There is a fixed sequence of muscle activation from proximal muscles to distal muscles. The order is preserved in all movements. For slow movements, it is shown that a constant joint stiffness is necessary to produce a smooth movement with a bell-shaped velocity. Scaled movements can be reproduced by varying the constraints on the maximal level of excitation signals according to the speed of movement. When the inertial parameters of the arm are altered, movement trajectories can be kept invariant by adjusting the pulse height values, showing the ability to adapt to load changes. These results agree with a wide range of experimental observations on human voluntary movements. Received: 4 December 1995 / Accepted in revised form: 17 September 1996     

Role of motor cortex in coordinating multi-joint movements: is it time for a new paradigm?

Reaching movements to spatial targets require motor patterns at the shoulder to be coordinated carefully with those at the elbow to smoothly move the hand through space. While the motor cortex is involved in this volitional task, considerable debate remains about how this cortical region participates in planning and controlling movement. This article reviews two opposing interpretations of motor cortical function during multi-joint movements. On the one hand, studies performed predominantly on single-joint movement generally support the notion that motor cortical activity is intimately involved in generating motor patterns at a given joint. In contrast, studies on reaching demonstrate correlations between motor cortical activity and features of movement related to the hand, suggesting that the motor cortex may be involved in more global features of the task. Although this latter paradigm involves a multi-joint motor task in which neural activity is correlated with features of movement related to the hand, this neural activity is also correlated to other movement variables. Therefore it is difficult to assess if and how the motor cortex contributes to the coordination of motor patterns at different joints. In particular, present paradigms cannot assess whether motor cortical activity contributes to the control of one joint or multiple joints during whole-arm tasks. The final point discussed in this article is the development of a new experimental device (KINARM) that can both monitor and manipulate the mechanics of the shoulder and elbow independently during multi-joint motor tasks. It is hoped that this new device will provide a new approach for examining how the motor cortex is involved in motor coordination.     

The contribution of the wrist, elbow and shoulder joints to single-finger tapping

We aimed to determine the role of the wrist, elbow and shoulder joints to single-finger tapping. Six human subjects tapped with their index finger at a rate of 3 taps/s on a keyswitch across five conditions, one freestyle (FS) and four instructed tapping strategies. The four instructed conditions were to tap on a keyswitch using the finger joint only (FO), the wrist joint only (WO), the elbow joint only (EO), and the shoulder joint only (SO). A single-axis force plate measured the fingertip force. An infra-red active-marker three-dimensional motion analysis system measured the movement of the fingertip, hand, forearm, upper arm and trunk. Inverse dynamics estimated joint torques for the metacarpal-phalangeal (MCP), wrist, elbow, and shoulder joints. For FS tapping 27%, 56%, and 18% of the vertical fingertip movement were a result of flexion of the MCP joint and wrist joint and extension of the elbow joint, respectively. During the FS movements the net joint powers between the MCP, wrist and elbow were positively correlated (correlation coefficients between 0.46 and 0.76) suggesting synergistic efforts. For the instructed tapping strategies (FO, WO, EO, and SO), correlations decreased to values below 0.35 suggesting relatively independent control of the different joints. For FS tapping, the kinematic and kinetic data indicate that the wrist and elbow contribute significantly, working in synergy with the finger joints to create the fingertip tapping task.     

Effect of axis alignment on in vivo shoulder kinematics

Background. To describe 3D shoulder joint movements, the International Society of Biomechanics (ISB) recommends using segment coordinate systems (SCSs) on the humerus, scapula and thorax, and joint coordinate systems (JCSs) on the shoulder. However, one of the remaining problems is how to define the zero angles when the arm is in an initial reference position. The aim of this paper is to compare various methods of determining the JCSs of the shoulder that make it possible to define the zero angles of the arm in the resting position.

Methods. Able-bodied subjects performed elevation movements in the scapular plane, specifically neutral, internal and external rotations of the humerus. The initial humerus position (at the beginning of the arm movement) and range of motion were analysed for the purpose of clinical interpretation of arm attitude and movement. The following four different JCSs were explored: (1) the standard JCS, defined as recommended by the ISB, (2) a first aligned JCS, where the humerus SCS is initially aligned with the scapula SCS, (3) a second aligned JCS, where the opposite operation is performed and 4) a third aligned JCS, where both the humerus and the scapular SCS are initially aligned with the thorax SCS.

Findings. The second aligned JCS was the only method that did not produce any exaggerated range of movement in either anatomical plane.

Interpretation. Mathematical JCS alignment allows clearer clinical interpretation of arm attitude and movement.     

Electrical activity of muscles during disturbances in the trajectory of single-joint motion in the arm

Horizontal "extension-flexion" movement of human arm at the elbow joint was studied. Humans performed movements in accordance with the instruction to ignore sudden trajectory changes that arose by the arm during lengthening of different elastic rods with friction. It was shown by multi-dimension regression analysis that the electric activity of biceps and triceps muscles of the shoulder was correlated first of all with the work of load and then with loading force, muscle work and other mechanic and myographic characteristics. One could suppose that information on energy quantities was contained in afferentation signals of the nervous system to correct the program commands on the spinal level. The purpose of correction was to preserve constant quantity of energy expenditure, as well as accuracy and duration of movement.